CSIRO PUBLISHING Australian Systematic Botany, 2019, 32, 555–563 https://doi.org/10.1071/SB19011

Resolving nomenclatural ambiguity in South American (Leguminosae, Papilionoideae, ), including the description of a new species

R. T. de Queiroz A,F, T. M. de Moura B,C, R. E. Gereau C, G. P. Lewis D and A. M. G. de Azevedo Tozzi E

ADepartamento de Sistemática e Ecologia, Centro de Ciências Exatas da Natureza, Universidade Federal da Paraíba (UFPB), Cidade Universitária, João Pessoa, PB, 58051-090, . BDepartamento Ciências Biológicas, Instituto Federal Goiano (IF Goiano), Rodovia Geraldo Silva Nascimento, quilômetro 2.5, Urutaí, GO, 75790-000, Brazil. CMissouri Botanical Garden, 4344 Shaw Boulevard, Saint Louis, MO 63110, USA. DComparative Plantand Fungal Biology Department,Royal BotanicGardens, Kew, Richmond, Surrey,TW9 3AB, UK. EDepartamento de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas (UNICAMP), Rua Monteiro Lobato 255, Cidade Universitária Zeferino Vaz, Barão Geraldo, Campinas, SP, 13083-862, Brazil. FCorresponding author. Email: [email protected]

Abstract. Taxonomic studies of Tephrosia Pers. (Leguminosae, Papilionoideae, Millettieae) in have highlighted the need to resolve some nomenclatural issues. Five new synonyms are proposed and a new species is described. Nine lectotypes of accepted names and synonyms, and one neotype, are here designated. An identification key to the taxa occurring in South America is also presented.

Additional keywords: , lectotypification, synonymy, systematics, .

Received 20 February 2019, accepted 31 July 2019, published online 7 October 2019

Introduction T. egregia Sandwith, T. fertilis R.T.Queiroz & A.M.G. Tephrosia Pers. (Leguminosae–Papilionoideae) comprises Azevedo, T. guaranitica Chodat & Hassl., T. hassleri Chodat, ~350 pantropically distributed species, occurring mainly in T. macbrideana R.T.Queiroz, G.P.Lewis & A.M.G.Azevedo, seasonally dry tropical woodlands, bushlands, thickets and T. marginata Hassl., T. noctiflora Bojer ex Baker, T. purpurea grasslands, often in open and disturbed sandy or rocky areas (L.) Pers. subsp. purpurea, T. senna Kunth, T. sessiliflora (Poir.) (Schrire 2005). It comprises two subgenera, namely, Tephrosia Hassl., and the new species described here, T. chaquenha subgenus Barbistyla Brummitt and Tephrosia subgenus R.T.Queiroz & A.M.G.Azevedo). Tephrosia. Tephrosia subg. Barbistyla has a pubescent style The species of Tephrosia in South America are shrubs and a glabrous stigma, whereas T. subg. Tephrosia has a or subshrubs with imparipinnate, multifoliolate, trifoliolate or pubescent stigma (Brummitt 1980). According to Schrire unifoliolate leaves. Inflorescences are terminal, axillary or (2005), the highest diversity for this is in and leaf-opposed pseudoracemes; the calyx is campanulate; the Madagascar (~170 species), (~90 spp.), Central and corolla is red, pinkish, purple, yellow, or white; and the ovary Tropical North America (~45 spp.) and Asia (~40 spp.). In South has 2–13 ovules. The fruit is a typical legume (Queiroz 2012). America, 18 taxa are recognised (Queiroz 2012). Bentham (1862) and Burkart (1952) noted that Tephrosia can be Most treatments of the species of Tephrosia in South easily recognised by its linear, elliptical, oblong or obovate America are in regional floras, e.g. Argentina (Burkart 1952), leaflets with numerous, parallel, oblique secondary veins. Brazil (Bentham 1862; Queiroz and Tozzi 2009, 2011; Brazilian Several species of Tephrosia are reported to have economic Flora Group 2015), Paraguay (Hassler 1919), Peru (Macbride importance. According to Forbes (1948), ~22 species have been 1943), Suriname (Amshoff 1939), and Venezuela (Pittier 1944). recorded as fish-poison . Wood (1949) noted that many Queiroz (2012) produced the only complete account of this genus species of this genus produce rotenone, a substance used to in South America, where Tephrosia subgenus Barbistyla is produce insecticides that are poisonous to invertebrates but not to represented by four species ( DC., T. nitens most vertebrates, and Dzenda et al.(2007) reviewed the Benth., T. sinapou (Buc’hoz) A.Chev., and T. vogelii Hook.f.), ethnomedical and veterinary uses of T. vogelii, and mentioned and Tephrosia subg. Tephrosia comprises 14 taxa (T. adunca eight other species of Tephrosia with therapeutic properties. Benth., T. cinerea (L.) Pers., T. domingensis (Willd.) Pers., Compounds isolated from this genus include deguelin, rotenone,

Journal compilation CSIRO 2019 Open Access CC BY-NC-ND www.publish.csiro.au/journals/asb 556 Australian Systematic Botany R. T. de Queiroz et al.

tephrosin, quercetin and rutin, which are reported as 3. Branches smooth, indumentum greyish; lenticels present on the antimicrobial and potentially able to kill molluscs, fish, branches; stipules 7–10-veined; leaflets oblanceolate; insects and helminths (Dzenda et al. 2007). pseudoracemes laxly flowered ...... T. nitens During his revision of Tephrosia in South America, the first 3: Branches striate, indumentum rusty brown; lenticels absent on author analysed more than 3000 accessions in North American, the branches; stipules 5-veined; leaflets oblong, elliptic to Brazilian and European herbaria. Materials from the following narrowly elliptic; pseudoracemes with flowers congested...... 4 herbaria were studied: B, BAB, BHCB, BM, BR, C, CEN, 4. Leaflets elliptic to narrowly elliptic; bracts ovate, caducous; CGMS, COL, E, EAC, EAN, ESA, F, FHI, G, GH, GOET, calyx 4-lobed, calyx tube shorter than the lobes; standard 2.5–2.6 3.0–3.2 cm, indumentum present on its abaxial HAL, HAS, HB, HEPH, HNBU, HRB, HRCB, HRJ, HST, – HUEFS, HUFMS, HUFU, IAC, IBGE, ICN, INPA, IPA, JPB, surface; legumes 10 20 mm wide, lanate ...... T. vogelii fl K,L,LP,M,MA,MBM,MG,MICH,MO,MOSS,NX,OUPR, 4: Lea ets narrowly oblong to oblong; bracts subulate to narrowly triangular, persistent; calyx with 5 lobes, calyx OXF, NY, P, PH, PAMG, PEUFR, RB, S, SI, SING, SP, SPF, tube longer than the lobes; standard 1.1–2.3 1–2.4 cm, SPFR, TCD, TEPB, UB, UC, UEC, UFG, UFMT, UFRJ, UFRN, indumentum absent on its abaxial surface; legumes 3–10 mm UFP, UPCB, U, US, W and WU (codes follow Index Herbariorum, wide, strigose to sparsely velutinous ...... 5 ’ New York Botanical Garden s Virtual Herbarium, see http:// 5. Leaves with 13–21 leaflets; these acute at the apex, and sweetgum.nybg.org/science/ih, accessed 22 February 2017). abaxially pubescent; inflorescence axis always longer This work has brought several nomenclatural issues to light. than the leaf length; pseudoraceme with up to As a result, we are here describing one new species, proposing 50 flowers; pedicel 0.8–1.5 cm long; calyx lobes nine lectotypes, one neotype, and five new synonyms. In rounded at apex; style apex straight; fruits laterally addition, an identification key to the taxa occurring in South flattened, 0.8–1 cm wide ...... T. candida America is presented. 5: Leaves with 27–39 leaflets; these retuse at the apex, and pubescent on both surfaces; inflorescence axis shorter than the leaf length; pseudoraceme with more than 50 Nomenclatural novelties flowers; pedicel 0.3–0.6 cm long; calyx lobes acute Tephrosia Pers., Syn. pl. 2(2): 328 (1807), nom. cons. at apex; style apex curved; fruits cylindrical, 4–5mm wide ...... T. sinapou Type: Tephrosia villosa (L.) Pers. 2: Branches slender; strigilose, hirsute, sericeous, sparsely sericeous, Cracca L., Sp. pl. 2: 752 (1753), nom. rej., non Hill (1756), nec Medik. pilose, or glabrescent; style and stigma glabrous; aril absent on (1787), nec Benth. (1853). seed ...... 6 Type: Cracca virginiana L. 6. Branches with a greyish indumentum or glabrescent ...... 7 Colinil Adans., Fam. Pl. 2: 327 (1763). 7. Branches sparsely sericeous to glabrescent; calyx lobes the Type: none. same length as the calyx tube ...... 8 fl – Erebinthus Mitch., Diss. Brev. Bot. Zool. 32 (1769), nom. rej. 8. Pseudoraceme laxly owered; calyx 4 5 mm long; corolla with dots; wings narrowly elliptic; leaflets rounded to Type: Tephrosia spicata (Walter) Torr. & A. Gray (fide acute at apex ...... T. domingensis C.Wood, Rhodora 51: 292 (1949)). 8: Pseudoraceme with flowers congested; calyx 5–7 mm long; Needhamia Scop., Intr. Hist. Nat. 310 (1777), nom. rej. corolla without dots; wings elliptic, falcate or obovate; fl Type: Vicia littoralis Jacq. lea ets retuse at apex ...... 9 9. Rachis 5 mm long; bracts over 3 mm long; wings Reineria Moench, Suppl. Meth. 44 (1802), nom. rej. 3-veined, claw over mm long; leaflets inserted Type: Reineria reflexa Moench. 1.5–2.2 cm apart, secondary veins salient on the Kiesera Reinw. ex Blume, Catalogus, 93 (1823). abaxial surface ...... T. senna – Type: Kiesera sericea Reinw. 9: Rachis 10 80 mm long; bracts up to 3 mm long; wings 2-veined, claw under 3 mm long; leaflets inserted Xiphocarpus C.Presl, Symb. Bot. 1: 13 (1830). 0.6–1 cm apart, secondary veins not salient on the Type: Xiphocarpus martinicensis C.Presl. abaxial surface...... T. purpurea subsp. Apodynomene E.Mey., Comm. Pl. Afr. Austr. 111 (1836). purpurea 7: Branches with a dense indumentum [except in T. chaquenha]; Type: Apodynomene grandiflora (L’Hér. ex Aiton) E.Mey. calyx lobes twice the length of the calyx tube ...... 10 Macronyx Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 2: 35 (1850). 10. Branches hirsute; leaflets over 4.7 cm long, tomentose Type: Macronyx strigosus Dalzell. abaxially; pseudoraceme congested; calyx lobes 9–14 mm long; standard 1.3–1.5 cm long; keel petals 1.3 cm long; staminal tube 1.1 cm long; fruit laterally Key to species of Tephrosia in South America compressed, hirsute ...... T. hassleri 10: Branches sericeous, pilose, tomentose, sparsely sericeous to 1. Leaves 3-foliolate, rarely 5-foliolate; rachis <2 cm long; corolla yellow; glabrescent; leaflets up to 4.5 cm long, sericeous abaxially; flowers in pairs on the inflorescence; calyx not persistent in fruit pseudoraceme laxly flowered; calyx lobes 1–7 mm long; ...... T. sessiliflora standard 0.5–1.3 cm long; keel petals 0.6–1.1 cm long; 1: Leaves rarely 3-foliolate, usually 5-foliolate; rachis >2 cm long; corolla staminal tube 0.6–1 cm long; fruit cylindrical, sericeous to white, purple or pink; inflorescence a pseudoraceme; calyx usually tomentose ...... 11 persistent in fruit ...... 2 11. Leaflets retuse at apex; calyx tube over 2 mm long, 2. Branches thick, strigose to densely strigose; hairs present on the style pubescent to tomentose, lobes 4–7 mm long; seeds but absent at the base of the stigma; aril present on the seed ...... 3 square ...... T. egregia Nomenclatural novelties in Tephrosia from South America Australian Systematic Botany 557

11: Leaflets acute to rounded at apex; calyx tube under Tephrosia penicillata Benth., Ann. Nat. Hist. 3: 431 (1839). 2 mm long, sericeous or hirsute, lobes 1–4mmlong; seeds oval to oblong ...... 12 Type: Guyana [British Guiana], s. dat., R.Schomburgk 678 (holo: K 502605; iso: BR 5792504, F 59923F, F 360659F, 12. Leaflets narrowly oblanceolate to oblanceolate or obovate; stipules 0.5–1 mm wide; bracts 1/2 the G 367963, G 367977, G 367990, NY 33832, TCD 4328, TCD pedicel length; standard 0.6–1 cm long ...... 4329)...... T. cinerea Tephrosia rufescens Benth., Linnaea 22: 513 (1849); Cracca villosa var. fl 12: Lea ets narrowly elliptic, elliptic, narrowly rufescens (Benth.) Kuntze, Revis. Gen. Pl. 1: 174 (1891); Tephrosia oblanceolate, or linear; stipules 0.5 mm wide; adunca var. rufescens (Benth.) Hassl., Repert. Spec. Nov. Regni Veg. 16: bracts approximately equally the length of the 165 (1919), syn. nov. pedicel; standard 1.2–1.3 cm long ...... 13 13. Pseudoraceme 20–50 cm long; an erect Type: Brazil. Minas Gerais, 1848, A.F.Regnell 78 (lecto, here subshrub; indumentum strigilose; stipules designated: K 502608; isolecto: R 64353, R 64353a, M (not 8–9 mm long; flowers 10–12 mm long seen), MO 1737286 (image seen). Remaining syn: Brazil, Nov...... T. macbrideana 1833, Riedel 1565 (P 2949532). Probable syn: Brazil., Minas 13: Pseudoraceme up to 13 cm long; plant a Gerais. Caxoeira do Campos, 1839, P.Claussen & B.Delessert 4 decumbent subshrub; indumentum (K 502607). sericeous; stipules 2–7 mm long; flowers 15–15.5 mm long ...... T. chaquenha Tephrosia adunca var. intermedia Chodat & Hassl., Bull. Herb. Boissier 6: Branches with a rusty brown indumentum ...... 14 sér. 2, 4: 839 (1904). 14. Plant an erect subshrub; adaxial surface of the leaflets glabrous; Type: Paraguay, Canindeyú, Yerbales. In regione Yerbalium calyx gibbous; ovary up to 6 mm long; fruit villous; seeds with a de Maracayú Paraguaria euro-astra, Serra de Maracayú, fl rugose testa, ochre-coloured ...... T. nocti ora E.Hassler 4643 (lecto, here designated: G 448507; isolecto: fl 14: Plant a decumbent subshrub; adaxial surface of the lea ets BM 538080, G 448505, G 448506, G 448506a, G 448507-a, sparsely sericeous, pilose, setose, or glabrescent; calyx G 448507-b, G 448507-c, G 448507-days, G 448508, G 448508- campanulate; ovary over 6 mm long; fruit pubescent, a, K 502603, UC 934893). sericeous or tomentose; seeds with a smooth testa, brown ...... 15 Tephrosia adunca f. glabrior Chodat & Hassl., Bull. Herb. Boissier, 15. Branches hirsute or pubescent; stipules linear; leaflets sér. 2, 4: 839 (1904). 2.2–2.8 cm wide; more than 25 pairs of veins per leaflet ...... T. guaranitica Type: Paraguay. In dumeto pr Bellavista in regione cursus fl – 15: Branches pilose or sericeous; stipules narrowly superioris uminis Apa, Oct. 1901 1902, E.Hassler 7843 (holo: triangular; leaflets up to 2 cm wide; less than 25 pairs G 448504). of veins per leaflet ...... 16 Tephrosia rufescens var. paraguayensis Ulbr., Repert. Spec. Nov. Regni fl 16. Lea ets linear to narrowly elliptic, never Veg. 2: 12 (1906); Tephrosia adunca f. paraguayensis (Ulbr.) Hassl., oblanceolate; marginal vein prominent; standard Repert. Spec. Nov. Regni Veg. 16: 165 (1919). 1.4 1.8 cm, transversally elliptic; wings 1.5–1.6 cm long; keel petals 1.1 cm long; ovary 1 cm Type: Paraguay, 20–25 Feb. 1903, K.Fiebrig 903 (lecto, here long ...... T. marginata designated: G 400050; isolecto: K 502609, M 233384). 16: Leaflets oblanceolate, narrowly oblong, elliptic, or narrowly elliptic; marginal vein not prominent; Tephrosia adunca var. subglabrata Hassl., Repert. Spec. Nov. Regni standard 0.7–1.2 0.8–1.4 cm, elliptic, widely Veg. 16: 165 (1919). – depressed-ovate to oblong; wings 0.8 1.1 cm Type: Paraguay. Zwischen Rio Apa und Rio Aquidaban, Nov. – – long; keel petals 0.6 1.0 cm long; ovary 0.7 0.9 cm 1908–1909, K.Fiebrig 4229 (lecto, here designated: G 448503; long ...... 17 isolecto: G 448503-a, G 440502, G 440502-a, G 440502-b, 17. Pedicel 1.5 shorter than the bracts; standard G 440502-c. Remaining syn: Paraguay. Cordillera de Altos, broadly depressed-ovate, standard claw 1.2 mm long; keel petals 10 mm long ...... 1903, K. Fiebrig 938 G 448500, G 448500-a, G 448501, ...... T. fertilis G 448501-a). 17: Pedicel twice the bract length; standard elliptic, Tephrosia adunca f. pseudomarginata Hassl., Repert. Spec. Nov. Regni broadly obovate or oblong, standard claw Veg. 16: 165 (1919). 1.7–4 mm long; keel petals 6–9 mm long ...... T. adunca Type: Paraguay. Prope Caaguazú in campis combustis, 1905, E.Hassler 8908 (holo: G 448441).

1. Tephrosia adunca Benth., Ann. Nat. Notes Hist. 3: 432 (1839) Tephrosia adunca has a wide geographical distribution Type: Brazil. Minas Gerais: in desertorum pascuis ad S. Isabel, (Argentina, Boliva, Brazil, the Guianas, Venezuela and s. dat., Pohl s.n. (lecto, here designated: K 502599; isolecto: NY Uruguay) and presents a large morphological variation. It is 623410). Cracca adunca (Benth.) Kuntze, Revis. Gen. Pl. 1: 174 morphologically similar to T. fertilis, but the latter can be (1891). recognised by its thick, curved branches (v. straight and 558 Australian Systematic Botany R. T. de Queiroz et al. thinner branches in T. adunca). The indumentum of T. adunca is duplicates, G 448502, G 448502-a, and G 448503 have flowers always rust-coloured. The leaf has a curved rachis when the and fruits and we, here, designate K.Fiebrig 4229 (G 448503) as leaflets are closed, and the leaflets are variable in form and the lectotype of Tephrosia adunca var. subglabrata Hassl. In the covered by a sparse to medium sericeous indumentum. A protologue of T. rufescens var. paraguayensis, Ulbrich (1906¸ diagnostic character of the species is that the pedicel is p. 165) cited ‘K. Fiebrig, Plantae Paraguayenses no. 903’. approximately twice the length of the bract. Ulbrich’s herbarium is housed at B (partly extant; Stafleu and The protologue of T. adunca cites a syntype collection: ‘Dr Cowan 1976). We did not locate any specimens that could Pohl in the desert pastures of S. Isabel’, and a second gathering be considered original material of Tephrosia adunca var. from Minas Gerais. We have located only two specimens paraguayensis Hassl. at B, but three specimens were located collected by Dr Pohl from S. Isabel (K 502599 and NY at G, M, and K. The specimen located at M (M 233384) is sterile. 623410). As the specimen at K includes an original Pohl Both K 502609 and G 400050 have flowers and fruits, in label, we are designating it as the lectotype of Tephrosia accordance with the protologue. Because G 400050 is the adunca Benth. most informative specimen (it has mature fruits, whereas the In the protologue of Tephrosia rufescens, Bentham (1862, other specimen has immature fruits), we are designating this as p. 513) cited three syntypes: ‘Ad Caldas prov. Minas Geräes the lectotype of Tephrosia rufescens var. paraguayensis Ulbr. A. Regnell. ser. 2. n.78; Caxoeiras do Campos, Claussen; ad San Paulo, Riedel’. According to Stafleu and Cowan (1976), George Bentham’s herbarium and types are housed at K, where we found two collections: Brazil, Minas Gerais, Caxoeira dos Campos, 2. Tephrosia chaquenha R.T.Queiroz & 1839, P.Claussen & B.Delessert 4 (K 502607) and Brazil, Minas A.M.G.Azevedo, sp. nov. Gerais, ad Caldas, 1848, A.F.Regnell 78 (K 502608). Both Type: Paraguay, Ciges Couchées, Villa Rica, sur les collines specimens are on the same sheet. An additional specimen, inculte, 6 Oct. 1874, B.Balansa 1539 (holo: BM 538072; iso: BR A.F.Regnell II-78 (K 858670, M 0233383), was also located; 5020184238698) (Fig. 1, 2). however, the label data indicate that this specimen was collected in Uberaba municipality, instead of Caldas, as cited in the Tephrosia cinerea f. pseudoadunca Hassl., Repert. Spec. Nov. Regni protologue. Duplicates of Regnell 78 collected at Minas Veg. 16: 166 (1919). Gerais, Caldas, were located at R (R 64353 and R 64353a). A probable syntype, L.Riedel 1565, is housed at P (P 2949532). Because Bentham cited the number 78 for the Regnell collection G in the protologue, and did not cite either the co-collector B. Delessert or the number 4 for the Claussen collection (therefore, we are not sure whether the Claussen specimen is indeed even a syntype), we are here designating K 502608 as the lectotype of Tephrosia rufescens Benth. H In the protologue of Tephrosia adunca var. intermedia,

Chodat and Hassler (1904, p. 834) cited ‘in campo pr. flumen 5 mm Jejui guazu, Sept., no. 4643’. Ten duplicates of Hassler 4643 were found at G. The following five of these specimens are I annotated as the holotype: G 448507, G 448507-a, G 448507-b, G 448507-c, G 448507-d; and the other five as isotypes: G 448505, G 448506, G 448506-b, G 448508-a, G 448508-b. All the specimens fit the protologue and are eligible for selection as the lectotype. Five of these sheets carry a specimen label (G 448505, G 448506, G 448507, G 448507-d and G 448508) and are equally informative. We opted to follow the 1 cm G labels, which annotate G 448507 as the holotype, and we 5 mm designate thisas the lectotype of the nameT.adunca var. intermedia. E In the protologue of Tephrosia adunca var. subglabrata, Hassler (1919, p. 165) cited the following three syntypes: ‘ ’ 5 mm Paraguay: Hassler 7843; Fiebrig 938, 4229 . According to K A Stafleu and Cowan (1979), the largest set of Émile Hassler’s Paraguayan specimens is housed at G, the second largest set at K, and other major sets at B (no longer extant), P, S and W. Material 5 mm 5 mm 5 mm of all cited syntypes is held at G. To avoid confusion, material of 5 mm J L Hassler 7843 was excluded from consideration for lectotype DCB F selection, as it is type material of the name Tephrosia adunca f. Fig. 1. Tephrosia chaquenha. A. Branch. B–D. Distinct leaflets. E. Bud. glabrior Chodat & Hassl. Four duplicates of Fiebrig 938 and six F. Calyx. G. Standard. H. Wings. I. Keel petals, J. Gynoecium, K. Staminal of Fiebrig 4229 are held at G. Fiebrig 4229 has flowers and fruits, tube. L. Young fruit with persitent calyx. Voucher Schinini et al. 19043 IAC- whereas Fiebrig 938 has only flowers. Among the Fiebrig 4229 24649. Drawn by Kley Souza. Nomenclatural novelties in Tephrosia from South America Australian Systematic Botany 559

A 10 μm

μ B 10 m

Fig. 2. Scanning electron micrographs of the testa of two species of Tephrosia. A. T. chaquenha (simple-reticulate papillate testa). B. T. cinerea (simple-reticulate smooth testa).

Type: Paraguay, Gran Chaco, Loma Clavel, E.Hassler 2591 to narrowly oblanceolate or linear; base acute to cuneate, apex (holo: G 229579; iso: G 229578, G 229580, G 400028, MPU rounded to acute; upper surface glabrous, lower surface scattered 024206, P 02950680 US 00003906). sericeous; secondary veins 7–9 pairs; all petiolules 1–2 mm long, Haec species Tephrosiae cinereae (L.) Pers. similis, sed ab ea sericeous. Inflorescence a terminal or axillary pseudoraceme praecipue foliolis adaxialitre glabris, floribus longioribus in 2.5–13 cm long; peduncle 3–5 cm long, cylindrical, scattered partibus omnibus majoribus atque petalis organis secretoriis sericeous to glabrous; bracts 2–5 0.4–1 mm, narrowly praeditis distinguitur. triangular; pedicels 3–5 mm long. Flowers pink, 15–15.5 mm Decumbent subshrub up to 80 cm tall, indumentum sparsely long; calyx 5–6 mm long, campanulate, bilabiate, scattered grey sericeous to glabrous. Leaves imparipinnate, 5–19- sericeous, teeth 2.8–4 ~1 mm, narrowly triangular, foliolate; stipules 2–7 ~0.5 mm, with 1–3 veins, narrowly subulate; standard petal 12–13 12–15 mm, broad ovate, triangular to linear; petiole 3–20 mm long, sulcate; rachis retuse at apex, dorsal surface scattered sericeous to tomentose, 5–50 mm long, sulcate; petiole, stipule, and rachis scattered claw 2–2.5 1–1.5 mm; wings 12–14 6–7 mm, asymmetrically sericeous; leaflets 10–45 3–7 mm, elliptic or narrowly elliptic obovate, apex rounded, claw 2–2.5 0.3–0.5 mm; keel petals 560 Australian Systematic Botany R. T. de Queiroz et al.

9–10 2–5 mm, connate, cucullate, apex acute, secretory Specimens examined – – structures present, claw 2.5 2.8 0.2 0.3 mm; staminal ARGENTINA. Corrientes: Lavalle, 2 km N de Cerrito sobre ruta 152, – – fi tube 8 10 2.5 3 mm, free portion of laments 23 Nov. 1979, A.Schinini 19043, R.Vanni & G.Norrmann (fl., fr.) (IAC); 3–4 mm long; annular disk present at base of sessile ovary; entreRios: Colón,13 Nov. 1979,N.S.Troncoso 2675 (fl.)(SI); Colón,Palmar, ovary 7–9 mm long, oblong, indumentum sericeous, ovules a 13 km E de R 14, barrancas del rio Uruguay hasta el arenal, lat. 3200S, long 8–10; style ~6 mm long, laterally compressed, glabrous; stigma 58100W, 17 Jan. 1976, C.Romanczuk 23 (fl., fr.) (BAB). BRAZIL. Mato penicillate with translucent trichomes. Fruit alegume43–60 Grosso do Sul: Porto Murtinho, Fazenda das flores 214300100S, 575305700W, 3–4 mm, coriaceous, plane, linear, apex and base asymmetric, 15 Dec. 2009, R.T.Queiroz 1439 (fl.) (UEC). PARAGUAY. Paraguay Centralis, fl brown with scattered sericeous, greyish indumentum, and in regione lacus Ypacaray, Dec. 1913, E.Hassler 12410 ( .) (BM, G, SI); persistent calyx; seeds 7–10 2.8–3.8 1.8–2.2 mm, oblong, Zwischen Rio Apa und Rio Aquidaban, 1909, K.Fiebrig 4229 (G, K). ochraceous, testa speckled, hilum excentric, aril absent. 3. Tephrosia cinerea (L.) Pers., Syn. Pl. 2: 328 (1807) Phenology Galega cinerea L., Syst. Nat., 10th edn, 2: 1172 (1759). Cracca villosa var. cinerea (L.) Kuntze, Revis. Gen. Pl. 1: 173 (1891); Cracca cinerea Flowers and fruits are reported from November to January. (L.) Morong, Ann. New York Acad. Sci. 7: 79 (1892); Colinil cinereum (L.) C.H.Hitchc., Rep. (Annual) Missouri Bot. Gard. 4: 75 (1893). Conservation status Type: [no specimen data] (lecto: LINN HL-924.2, designated by Tephrosia chaquenha is currently recorded from Argentina, W.Fawcett and A.B.Rendle, Fl. Jamaica 4: 20 (1920)). Brazil and Paraguay. The extent of occurrence (EOO) for the Vicia littoralis Jacq., Enum. Syst. Pl. 27 (1760). Galega littoralis species is 235906.527 km2 (which equates to a conservation (Jacq.) L., Syst. Nat., 12th edn, 2: 497 (1767); Tephrosia littoralis (Jacq.) status of Least Concern, LC). Its area of occupancy (AOO) is Pers., Syn. Pl. 2: 328 (1807); Tephrosia cinerea var. littoralis (Jacq.) 24.000 km2 (which would give it an Endangered status, EN). We Benth. in C.F.P. von Martius (ed.), Fl. Bras. 15(1A): 48 (1859); Cracca have little information about the quality of the habitats where the littoralis(Jacq.) Rydb., in N.L.Brittonet al. (eds),N. Amer.Fl. 24(3):178 species grows in each country, and because of the current paucity (1923). of specimens and habitat data, we assess Tephrosia chaquenha to Type: [no specimen data] (lecto, here designated: LINN HL- be Data Deficient (DD) following IUCN 2017 criteria (IUCN 924.3). 2017). Tephrosia venustula Kunth., nov. gen. sp., 4th edn 6: 459 (1823).

Etymology Type: Guyana. Bordones, s. dat., F.Humboldt & A.Bonpland 420 (holo: P 660150). The species is named in reference to its distribution in chaco vegetation (in Brazil and Paraguay). The habitat of this species in Orobus domingensis Spreng., Syst. Veg., 16th edn, 3: 261 (1826). Argentina is presently unknown. Type: Republica Dominicana. Santo Domingo, En terreno abandonado, común, cerca del Jardim Botânico, Santo Domingo, Notes 20 Sept 1973, A.H.Liogier 20188 (neo, here designated: NY Tephrosia chaquenha morphologically resembles T. cinerea and 1652524). shares the same habit, grey-coloured indumentum and similar linear fruits. The species can be distinguished by the characters Tephrosia procumbens Macfady., Fl. Jamaica 1: 256 (1837). presented in Table 1. Moreover, another noteworthy character is Type citation: ‘Galega No. 2, Browne, 289.’ that Tephrosia cinerea presents a reticulate-smooth testa, whereas it is reticulate-papillate in T. chaquenha (Fig. 2). Type: not seen.

Table 1. Characters used to distinguish Tephrosia cinerea from T. chaquenha SEM, Scanning Electron Microscopy

Character T. cinerea T. chaquenha Indument Pilose Scattered sericeous or glabrous Leaflets Oblanceolate or narrowly Elliptic or narrowly elliptic to narrowly oblanceolate oblanceolate or linear Upper surface of leaflets Sericeous Glabrous Inflorescence Solitary, axillary, flower, Terminal pseudoraceme, Flower length 0.8–15 mm 15–15.5 mm Calyx length 0.5–5mm 5–6mm Calyx symmetry Actinomorphic Bilabiate Standard petal length 6–10 mm 12–13 mm Petal secretory structures Absent Present Standard claw length 1–2 0.2–0.3 mm 2–2.5 1–1.5 mm Wing petal claw length 0.9–1.1 0.2–0.4 mm 2–2.5 0.3–0.5 mm Keel petal claw length 1.2–2 mm 2.5–2.8 mm Testa of seed at SEM (Fig. 2B) Simple-reticulate, smooth Simple-reticulate, papillate Nomenclatural novelties in Tephrosia from South America Australian Systematic Botany 561

Tephrosia gynothrix Miq., Linnaea 18: 29 (1844). Tephrosia tenella A.Gray, Smithsonian Contr. Knowl. 5(6):36 (1853). Cracca tenella (A.Gray) Rose, Contr. USA Natl. Herb. 12(7): 271 Type: Suriname. In plantatione de Zwarigheid, Vredenburger (1909). Kreek, Oct. 1842, H.C.Focke 666 (holo: U 3650). Type: United States. San Pedro: Sonora, 8 Sep. 1851, Tephrosia scopulorum Brandegee, Univ. Calif. Publ. Bot. 6: 181 (1915). C.Wright 966, (lecto, here designated: GH 107070; isolecto: Type: Mexico. San Geronimo, Oaxaca, July 1914, GH 63573, GH 63572, K 848919, K 848920, MO 217373, MO C.A.Purpus 7151 (holo: UC 175091; iso: F F0059920F, GH 277489, NY 33799, PH 30249, US 3979). (not seen), MO 125154, NY 33816, US 3953). Tephrosia leptostachya var. leptophylla Benth. in C.F.P. von Martius Tephrosia cinerea var. typica Hassl., Repert. Spec. Nov. Regni Veg. 16: (ed.), Fl. Bras. 15(1A): 49 (1862), syn. nov. 166 (1919), nom. inval. Type: Brazil. Goiás: Alegre, s. dat., Pohl s.n. (holo: K 921500).

Notes Cracca rusbyi Rydb. in N.L.Britton et al. (eds), N. Amer. Fl. 24(3): 181 (1923). This species has a subshrubby habit, with decumbent and thin branches. In general, the branches, leaves and fruits are covered Type: Mexico. Santa Catarina, Oaxaca, 14 July 1910 by grey indumentum. All material from Caatinga vegetation has H.H.Rusby 69 (holo: NY 6596). axillary fascicles of flowers with long, slender pedicels. The length of the pedicel is a character that facilitates the recognition of this species. Furthermore, its fruits are usually very long and Notes arched. Owing to the slenderness of the pedicel, the fruits are always orientated downward. The calyx lobes are strongly The name Tephrosia domingensis dropped into disuse after subulate, especially in fruit. The species is morphologically Bentham (1862) treated it as a synonym of T. leptostachya similar in appearance to T. chaquenha, but is easily DC. and published T. leptostachya var. leptophylla Benth., on distinguished by the above suite of characters the basis of material of T. domingensis. After that, all South The protologue of Vicia littoralis Jacq. (1760) cited neither American collections were treated as T. leptostachya. After specimens nor illustrations. Because Jacquin’s specimens are reviewing the type specimen of T. domingensis, we consider known to be in numerous herbaria (D’Arcy 1970; Stafleu and that the South American material treated as T. leptostachya is Cowan 1979), searches for original material were undertaken at better considered as a separate species, T. domingensis. B, BM, CGE, H, JE, LINN, LIV, OXF, UPS and W. We located Tephrosia domingensis is recognised by its decumbent habit, only one specimen (LINN HL-924.3) annotated ‘Vicia littoralis but in some situations it may become ascendant. Its branches are mea Jacquin’ in Jacquin’s handwriting (Burdet 1979), and, slender, strongly lignified, sometimes wine red in colour, and therefore, it is likely that it was seen by Jacquin. Accordingly, glabrescent. The leaves have very narrow and glabrescent leaflets we designate this specimen as the lectotype of Vicia litoralis with the apex usually acute, but occasionally round. The Jacq. An additional four specimens held at LINN (LINN pseudoracemes are long with few flowers. Their legumes are HS-1214.10, LINN HS-1214.11, LINN HS-1214.12, and plane, with the edges of the valves prominent and with a persistent LINN HS-1214.14) lack locality or date information, and each calyx; the calyx lobes are very delicate. The seeds are usually is annotated as Galega cinerea L. These specimens are unlikely flattened, and become dark and grey-speckled when old. to represent the original material of Vicia littoralis. The other In the protologue of Tephrosia tenella, Gray (1853) stated specimen located in one of the herbaria that may hold Jacquin ‘...near the San Pedro, Sonora; Sept. (966)’. Stafleu and Cowan material is W 29662, also lacking location or date, with the name (1976) stated that the herbarium, types, original manuscripts, Galega littoralis L. written on it. archives and correspondence of Asa Gray are at GH. Three Orobus domingensis Spreng. was described on the basis of a specimens of C.Wright 966 were located at GH (GH 63572, GH Carlo Bertero collection from Hispaniola in Santo Domingo. The 63573, and GH 107070). The protologue describes flowers and protologue provides neither a description nor an illustration. fruits, and all the three specimens fit the protologue. The Searches for original material of this name in B, BHU, BM, G, K, specimen GH 107070 has a label on it (certainly not prepared M, P and W were unsuccessful. Therefore, the specimen NY by Gray), identifying it as the holotype. However, on the same 1652524 is here proposed as a neotype for Orobus domingensis. sheet, there are two different collections, namely, GH 107070 (two plants on right half of sheet) and GH 263484 (two plants on 4. Tephrosia domingensis (Willd.) Pers., left half of sheet, with the collection number 929 on the label). Syn. Pl. 2(2): 330 (1807) Because GH 107070 is the most informative specimen, with both flowers and well-developed fruit, it is here formally Galega domingensis Willd., Sp. Pl., 4th edn, 3(2): 1249 (1802); Cracca designated as the lectotype of T. tenella A.Gray. domingensis (Willd.) Rydb., in N.L.Britton et al. (eds), N. Amer. Fl.24 (3): 181 (1923). Type: Dominica. ...St. Domingo..., s. dat., F.Bredemeyer s.n. (holo: B-W 13942-010). 5. Tephrosia marginata Hassl., Repert. Spec. Nov. Regni Veg. 16: 162 (1919), nom. nov. Tephrosia ascendens Macfady., Fl. Jamaica, 1: 257 (1837). Type: Paraguay. In valle fluminis Y – acá. in campo glareoso, pr. Type: Jamaica, s. dat., J.Macfadyen s.n. (holo: K 502616). Pirihebuy, Dec. 1900, E.Hassler 6721 (holo: G 400037). 562 Australian Systematic Botany R. T. de Queiroz et al.

Tephrosia nervosa Chodat & Hassl., Bull. Herb. Boisser, sér. 2, 4: 839 In the protologue of Tephrosia marginata var. (1904), nom. illeg., non Pers. (1807). Tephrosia adunca var. acutifolia pseudorufescens, Hassler (1919, p. 163) cited the syntypes Chodat & Hassl., Bull. Herb. Boissier, sér. 2, 4: 879 (1904), syn. nov. ‘Hassler 4346, 4381 id. in campis siccis Caaguazú Hassler ’ Type: Paraguay, Iter ad Yerabales montium Sierra de 9126 . Several specimens were located at BM, G, K, and Maracayu, in regione flumis Capibari, 1900, E.Hassler 4381 NY. Among the specimens at G (where the Hassler types are (lecto, here designated: G 400034; isolecto: G 400033, housed), G 400038 has only fruits, whereas the others have both fl G 400035, G 448509, G [no barcode accession number]. owers and fruits. Because the protologue describes fruits but does not mentionflowers,weare here designating G400038asthe Tephrosia marginata var. pseudorufescens Hassl., Repert. Spec. Nov. lectotype of Tephrosia marginata var. pseudo-rufescens Hassl. Regni Veg. 16: 163 (1919). Type: Paraguay. In campo Nandurucay Sierra de Maracayu, 6. Hook. f., Niger Fl. 296 (1849) 1898–1899, E.Hassler 4346 (lecto, here designated: G 400038; Cracca vogelii (Hook. f.) Kuntze, Revis. Gen. Pl. 1: 175 (1891). isolecto: G 400036, G 400044, NY 33830). Remaining syn: Type: Nigeria. Niger, s. dat., Vogel s.n. (holo: K 262618). Paraguay. In regione Yerbalium de Maracayú, E.Hassler 4381, BM 538077, G 400033, G 400034, G 400035, K 502595, Tephrosia megalantha Micheli, Bull. Soc. Roy. Bot. Belgique 36: 57 NY 33831; Paraguay, Prope Caaguazú in campis combustis, (1897), syn. nov. E.Hassler 9126, G 400026). Type: Democratic Republic of the Congo. Lusambo, s. dat., E.Laurent s.n. (lecto, here designated: BR 8947048). Tephrosia marginata var. cinerascens Hassl., Repert. Spec. Nov. Regni Veg. 16: 163 (1919), nom. inval. Tephrosia periculosa Baker, Bull. Misc. Inform. Kew. 1897 128–129: 258 (1897), syn. nov. Notes Type: Malawi. Kondowe to Karonga, s. dat., A.Whyte 324 Chodat and Hassler (1904) described this species as Tephrosia (holo: K 262825). nervosa on the basis of material collected in Paraguay. However, this name was an illegitimate later homonym, and Hassler (1919) Notes subsequently published the replacement name T. marginata and Tephrosia vogelii is a well-circumscribed species. In the used characters of the indumentum to describe two varieties, vegetative stage, its branches are strongly grooved. Both the namely, Tephrosia marginata var. cinerascens Hassl., nom. branches and the leaves are densely covered by strigose and inval. and T. marginata var. pseudorufescens Hassl. However, rufous trichomes. The species can be recognised more easily in the diagnostic characters used to separate the varieties are the reproductive state by its caducous bracts, a character that is tenuous, and these taxa are not accepted here. Tephrosia absent in other South American species. The flowers have a marginata var. cinerascens is invalid under ICN Art. 26.2 tomentose indumentum on the calyx lobes, and the standard has (Turland et al. 2018; known informally as ‘The Shenzhen indumentum on its dorsal surface; these characters are present Code’ or simply ‘the Code’, see http://www.iapt-taxon.org/ only in this species. The indumentum of the fruit is woolly, which nomen/main.php) because one of the cited syntypes (Hassler is another exclusive character of this introduced species in South 6721) is the type of T. nervosa (= T. marginata). America. This species morphologically resembles Tephrosia Tephrosia vogelii was introduced to South America probably chaquenha regarding the shape of the leaflets. However, because of its ornamental potential and ichthyotoxic properties. T. marginata is the only species with linear leaflets with a Most likely, the species originated in Central Africa. It is grown prominent marginal rib. Besides this, T. marginata has a in Africa for the extraction of insecticidal substances. In South ferruginous indumentum, long pseudoraceme (8–20 cm long), America, T. vogelii is infrequently encountered and only a small and wine red-coloured flowers, whereas T. chaquenha presents a number of collections exist in herbaria. grey indumentum, short pseudoraceme (2.5–13 cm long) and In the protologue of Tephrosia megalantha, M. Micheli pink flowers. (Leguminosaceae, cited in Chalon et al. 1897, pp. 53–77) In the protologue of Tephrosia adunca var. acutifolia, Chodat cited ‘Congo (Dewèvre n. 520)’ and ‘Lusambo (Laurent).’ and Hassler (1904) did not cite any collection. Nevertheless, According to Stafleu and Cowan (1981), the herbarium and Chodat and Hassler (1907, p. 826) published a correction to their types of Marc Micheli are housed at G, but no original original protologue citing E.Hassler 4381 as the type collection. material was located in this herbarium. One specimen of We are confident, therefore, that this is the original collection E.Laurent s.n. was located at BR. Therefore, we are designating analysed by the authors when describing their new taxon. Four BR-8947048 as the lectotype of Tephrosia megalantha Micheli. specimens under this number were located at G (G 400033, G 400033a, G 400034, G 400035, and G 448509). We Conflicts of interest are designating G 400034 as the lectotype of T. adunca var. acutifolia Chodat & Hassl., because this is the most informative The authors declare that they have no conflict of interest. specimen, having flowers and young fruits (other duplicates have only flowers). Moreover, the original description and the type Declaration of funding collection present no morphological characters that differentiate We thank CAPES and CNPq (processes 563550/2010-4 and the taxon from T. marginata; therefore, we are synonymising 201550/2010-5), and FAPESP (process 2010/52488-9) for T. adunca var. acutifolia under T. marginata. financial and logistical support to the first author. Nomenclatural novelties in Tephrosia from South America Australian Systematic Botany 563

Acknowledgements Subcommittee. (International Union for Conservation of Nature) Available at http://www.iucnredlist.org/documents/RedListGuidelines. We thank Dr Alain Chautems and Dr Jim Solomon for providing help while pdf [Verified 16 March 2019]. visiting G and MO respectively; we also thank Dr Laurent Gautier and Macbride JF (1943) Leguminosae. Field Museum of Natural History, Laurence Loze (at G) who kindly scanned and sent us the images of many Botanica Serbica 13,3–507. specimens, and Lorenzo Ramella (also at G) who provided type information; PittierH (1944)‘Leguminosasde VenezuelaI. Papilionaceae.’(Ministério da and the curators and staff at the herbaria cited in the text for their support and Agricultura y Cria, Elite: Caracas, Venezueala) forthe loanof specimens.Finally, the authorsthank theanonymous reviewers Queiroz RT (2012) Revisão taxonômica das espécies do gênero Tephrosia for their contribution and the editors, Dr Ashley Egan and Dr Daniel Murphy, Pers. 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